Are animal personality, body condition, physiology and structural size integrated? A comparison of species, populations and sexes, and the value of study replication.

A major question in behavioural ecology is why behaviour, physiology and morphology are often integrated into syndromes. In great tits, Parus major, for example, explorative males are larger (vs. smaller) and leaner (vs. heavier) compared to less explorative individuals. Unfortunately, considerable debate exists on whether patterns found in specific studies are replicable. This debate calls for study replication among species, populations and sexes. We measured behavioural (exploration), physiological (breathing rate) and morphological traits (body mass, tarsus length, wing length, bill length) in two species (great vs. blue tits Cyanistes caeruleus), two populations (Forstenrieder Park vs. Starnberg) and two sexes (males vs. females). We then tested whether the same pattern of integration characterized all unique combinations of these three biological categories (hereafter called datasets). We used a multi-year repeated measures set-up to estimate among-individual trait correlation matrices for each dataset. We then used structural equation modelling to test for size-dependent behaviour and physiology, size-corrected (i.e. size-independent) behaviour-physiology correlations and size-corrected body mass-dependent behaviour and physiology. Finally, we used meta-analyses to test which structural paths were generally (vs. conditionally) supported (vs. unsupported). We found general and consistent support for size-dependent physiology and size-corrected body mass-dependent physiology across datasets: faster breathers were smaller but heavier for their size. Unexpectedly, condition-dependent behaviour was not supported: explorative birds were neither leaner, nor was this relationship heterogeneous across datasets. All other hypothesized patterns were dataset-specific: the covariance between size and behaviour, and between behaviour and physiology differed in sign between datasets, and both were, on average, not supported. This heterogeneity was not explained by any of our moderators: species, population or sex. The specific pattern of size- and condition-dependent physiology reported for a unique combination of species, population, and sex, thus generally predicted those in others. Patterns of size- or condition-dependent behaviour (i.e. 'personality'), or behaviour-physiology syndromes reported in specific datasets, by contrast, did not. These findings call for studies revealing the ecological background of this variation and highlight the value of study replication to help understand whether patterns of phenotypic integration reported in one study can be generalized.

The association between personalities, alternative breeding strategies and reproductive success in dunnocks.

Although consistent between-individual differences in behaviour (i.e. animal personality) are ubiquitous in natural populations, relatively few studies have examined how personalities influence the formation of social relationships. Yet, behavioural characteristics of both sexes might be key when it comes to pair-bond formation, and cooperation with partners to successfully rear offspring. We here use a wild population of dunnocks (Prunella modularis) to first investigate whether individuals mate nonrandomly (i.e. assortative mating) with regard to four behavioural traits-flight-initiation distance (FID), provisioning, activity and vigilance-that differ in repeatability and have previously been associated with mating patterns and fitness in other species. Second, we test whether an individual's FID is associated with variability in the dunnocks' mating system (i.e. monogamous pairs vs. polygamous groups). Finally, we determine whether FID and provisioning of males and females associate with their reproductive success. We found no statistical support for assortative mating in FID between males and females. Interestingly, in polygamous groups, co-breeding males differed in their FIDs with dominant alpha males having significantly shorter FIDs compared with subordinate beta-males. Moreover, there was evidence for assortative mating in provisioning for alpha males and females in polygamous groups. We also found that male provisioning influenced reproductive success of both sexes, whereas female provisioning rates only positively correlated with her own but not their partner(s) reproductive output. Our results suggest that personality differences may have important implications for social relationships, the emergence of different mating patterns and ultimately reproductive success within populations.

Strong phenotypic trait correlations between mating partners do not result from assortative mating in wild great tits (Parus major).

There is considerable debate about the occurrence of assortative mating between phenotypic traits measured within natural populations. Meta-analyses have implied that assortative mating occurs generally in natural populations, but recent work indicates these conclusions largely result from biased data. Specifically, estimates of phenotypic correlations between mating partners do not solely result from nonrandom associations between individual-level traits of partners but also from other biological processes (joint phenotypic plasticity, indirect genetic effects), methodological practices (observer bias) and other unexplained residual correlations (e.g. correlated measurement error). This paper puts this critique to test. First, we estimated the overall phenotypic correlation between phenotypic traits of mating partners for a wild population of great tits. Second, we estimated various key variance components to reveal the extent to which phenotypic correlations between partners resulted from assortative mating, reversible plasticity, social partner effects and methodological practices. We performed our analyses for a range of phenotypic traits (body mass, breathing rate, exploration behaviour, wing and tarsus length) to derive general conclusions not hinging on the specifics of the traits involved. Our analyses support the conclusion that patterns of assortative mating exist at first glance but occur because of the biasing effects of correlated residuals likely caused by a combination of phenotypic responses to unknown environmental factors or measurement error-not because of intrinsic patterns of assortative mating.

Dunnock social status correlates with sperm speed, but fast sperm does not always equal high fitness.

Sperm competition theory predicts that males should modulate sperm investment according to their social status. Sperm speed, one proxy of sperm quality, also influences the outcome of sperm competition because fast sperm cells may fertilize eggs before slow sperm cells. We evaluated whether the social status of males predicted their sperm speed in a wild population of dunnocks (Prunella modularis). In addition to the traditional analysis of the average speed of sperm cells per sample, we also analysed subsamples of the fastest sperm cells per sample. In other words, we systematically evaluated the effects of including different numbers of the fastest sperm in our analyses, ranging from the 5-fastest sperm cells to the 100-fastest sperm cells in a sample. We further evaluated whether fitness, defined here as the number of chicks sired per male per breeding season, relates to the sperm speed in the same population. We found that males in monogamous pairings (i.e. low levels of sperm competition), produced the slowest sperm cells, whereas subordinate males in polyandrous male-male coalitions (i.e. high levels of sperm competition) produced the fastest sperm cells. This result was consistent regardless of the number of fastest sperm included in our analyses, but statistical support was conditional on the number of sperm cells included in the analysis. Interestingly, we found no significant relationship between fitness and sperm speed, which suggests that it is possible that the differential mating opportunities across social status levelled out any possible difference. Our study also suggests that it is important to identify biologically meaningful subsets of fastest sperm and cut-offs for inclusions for assessing sperm competition via sperm speed.

Dominance relationships and coalitionary aggression against conspecifics in female carrion crows.

Cooperation is a prevailing feature of many animal systems. Coalitionary aggression, where a group of individuals engages in coordinated behaviour to the detriment of conspecific targets, is a form of cooperation involving complex social interactions. To date, evidence has been dominated by studies in humans and other primates with a clear bias towards studies of male-male coalitions. We here characterize coalitionary aggression behaviour in a group of female carrion crows consisting of recruitment, coordinated chase, and attack. The individual of highest social rank liaised with the second most dominant individual to engage in coordinated chase and attack of a lower ranked crow on several occasions. Despite active intervention by the third most highly ranked individual opposing the offenders, the attack finally resulted in the death of the victim. All individuals were unrelated, of the same sex, and naïve to the behaviour excluding kinship, reproduction, and social learning as possible drivers. Instead, the coalition may reflect a strategy of the dominant individual to secure long-term social benefits. Overall, the study provides evidence that members of the crow family engage in coordinated alliances directed against conspecifics as a possible means to manipulate their social environment.

Personality-matching habitat choice, rather than behavioural plasticity, is a likely driver of a phenotype-environment covariance.

An emerging hypothesis of animal personality posits that animals choose the habitat that best fits their personality, and that the match between habitat and personality can facilitate population differentiation, and eventually speciation. However, behavioural plasticity and the adjustment of behaviours to new environments have been a classical explanation for such matching patterns. Using a population of dunnocks (Prunella modularis), we empirically tested whether personality or behavioural plasticity is responsible for the non-random distribution of shy and bold individuals in a heterogeneous environment. We found evidence for bold individuals settling in areas with high human disturbance, but also that birds became bolder with increasing age. Importantly, personality primarily determines the distribution of individuals, and behavioural adjustment over time contributes very little to the observed patterns. We cannot, however, exclude a possibility of very early behavioural plasticity (a type of developmental plasticity) shaping what we refer to as 'personality'. Nonetheless, our findings highlight the role personality plays in shaping population structure, lending support to the theory of personality-mediated speciation. Moreover, personality-matching habitat choice has important implications for population management and conservation.